Biological Psychology: A Brief History

…puerile, dull, dogmatic, absurd, foolish, and …the most frontless

piece of charlatanry that the age had produced, while the "cunning
craniologers" who were beginning to roam the country were seen as
quacks, empirics, manipulating impostors, and itinerant mountebanks
to be looked upon "as rather knaves than fools." (Cooter, 1984, p. 22)
 Phrenology's Modern Ghosts
The quotation above refers to phrenology, long regarded a
pseudoscientific fraud, but initiated by Franz Josef Gall, an eminent
Viennese physician and anatomist, who was the first to distinguish the white
and gray matter of the brain. He gathered masses of data relating "mental
and moral" attributes to the shape of people's heads. He proposed 26
faculties, or "personality organs," each complete and located in specific sites
in the brain. If we have a lot of a trait, such as facility with language, the
brain will be highly developed in the corresponding area and will cause the
skull to bulge at that point.
 The phrenologist, also called craniologist, zoonomist,
physiognomist, and other things could then analyze personality by
examining the surface of the skull.

Organ number two is Philoprogenitiveness, love of children, and is

located at the back of the head, since Gall believed this area to be
prominent in women and in apes, both of whom supposedly love children
more than do men. Organ number 22 is Individuality, located
immediately above the nose, since that is what seemed large in the face
of Michelangelo and small in Scottish people!

It may seem that phrenology is ridiculed today, but if you attend to

newspapers and television you will see that the general point of view
remains popular - we still appear to believe in mental motives and
powers (faculties) that are localized in specific brain areas.

Sometimes they are conceived as existing in their entirety in some

location - perhaps as "memory units" in the cerebral cortex. "Didn't
someone, Penfield, I think, revive memories by stimulating the cortex of
patients?" we think. “Scientists Discover Pleasure Centers in the Brain”
headlined a Montreal newspaper in the 1950s and variations on that
announcement have appeared frequently in newspapers and magazines
since.
You may also have heard of "aggression centers" in the brain. As recently
as the late 1960s and the 1970s many people were operated upon by
surgeons who believed that there were such centers (see Valenstein,
1971).

We hear also that there are "feeding and drinking centers" in the brain and
every decade at least two or three "satiety hormones" are reported, each
hoped to be the chemical that will turn off our hunger and make us slim.

Sometimes it is the "arousal" center that makes news or dopamine that

causes "awakenings" of Parkinsonism patients. In countless cases the
spirit of phrenology lives on in the localization of faculties in discrete brain
structures or in the amazing effects of a new hormone. The same applies
to the fruits of so-called "split brain" research.
 Richard Gregory noted that laterality theories, which posit
language functions in the left cerebral hemisphere and spatial
apprehension in the right, tell an old tale:
One might say that the current interest in 'cerebral dominance,' with
the left hemisphere of the cortex supposedly 'analytic' (responsible for
skills such as arithmetic and logical thinking) and the right hemisphere
'synthetic' or 'analogue' (responsible for intuitive and artistic skills), is
the dying kick of phrenology. (Gregory, 1987, p. 619-620)
In fact, if we add up all of the claims for "centers" of one kind or
another in the brain, we could list most of the motives and powers
proposed by the phrenologists. Almost a dozen separate functions
have been attributed to one brain structure, the hippocampus, and it
isn't even in the cerebral neocortex. Does that support phrenology?
 Localization of Function: Finding the Right Names for
Functions

Let us examine the evidence and briefly look at the history of

research in this area.

We will find that specific parts of the brain do serve specific

functions, as it seems they must, but it will also be clear that these
functions are often hard to name. That is, when we list functions
like attention, memory, learning, arousal, hunger, aggression, and
the like, our list seems not to correspond to the functions built in by
the architect of the nervous system.

Happily, recent research seeking the classic "centers" of the

phrenologists has brought a new understanding of the
characteristics of the nervous system. This allows a revision of the
list of centers, thus advancing our knowledge of psychology, as
well as of neuroanatomy and physiology.
 Nineteenth-Century Science
The Neural Impulse
Luigi Galvani
Alessandro Volta
The Rate of Neural Conduction
Hermann Helmholtz
Sensory and Motor Nerves
Sir Charles Bell
Francois Magendie
Reflex Action
Robert Whytt
Marshall Hall
Sherrington
Reflexes of the Brain
Ivan Sechenov
Specific Nerve Energies
Johannes Müller
Motor Cortex
Fritsch & Hitzig
Somatosensory cortex
Bartholomew
 Merzenich And A New Twist

In the 1980s M. M. Merzenich discovered that the

receptive fields in the somatosensory cortex of the postcentral
gyrus are surprisingly malleable.

For example, after amputation of a finger, the cortical

receptive field for that finger is "invaded" by the receptive fields
for the neighboring digits.

This totally-unexpected malleability of the sensory cortex –

the “map” of the body surface – changed our view of the cortex
as a set of fixed maps influenced by “association” cortex.
 Broca's "Language Center:" Lesson in Interpretation

The search for brain targets subserving specific functions appeared

to take a giant leap forward in 1861 when Pierre-Paul Broca discovered
what appeared to be the "speech center."

A patient described as otherwise normal was unable to articulate

language, though he could understand it. He was called "Tan," since
"tan-tan" were the only syllables he could utter. For years he had been
paralyzed on the right side, evidently due to a series of strokes (to which
we refer later).
He was referred to Broca for treatment of an infected bedsore and
soon died. Upon autopsy he was found to have a lesion in the left
hemisphere of the lower temporal lobe just forward of the temporal lobe
(the inferior posterior frontal lobe) - an area scarcely four centimeters
square. For a century this area was known as the speech center - its
function turned out to be not what it seemed to Broca.
 Reassessing Broca's Finding

Karl Pribram is an eminent neurosurgeon and psychologist who

believes that patients showing "Broca's aphasia" may suffer from more
general disabilities than the loss of the ability to speak. His
investigation shows that Broca did indeed find what he was expecting
to find.
Pribram (1971) wrote:

“Broca had been taught that language was a function of the

frontal lobes; his teachers derived their doctrine from the phrenologists
who had reasoned that man's high forehead and his linguistic ability
were two of his most distinguishing features; ergo they might well be
related. Broca reasoned that the only place where his aphasic
patient's lesion overlapped the frontal cortex was in the posterior
inferior portion. Hence Broca's area.

Broca's patient had suffered strokes that involved the middle

cerebral artery; this produces widespread damage, but it was the
damage to the frontal cortex, "Broca's area," that Broca noted. Is the
area the seat of speech?
Pribram suggests that it is not:
“Evidence against Broca's claim is simply that excision of
Broca's area in man's brain, and damage to this area, has been
inflicted without causing any severe linguistic disturbance...
Presumably therefore all of the lobotomies performed for
psychosurgical reasons (over 10,000) injured Broca's area to
some extent. Yet not a single report of aphasia due to lobotomy
occurred.” (Pribram, 1971)

In two catatonic patients who had not spoken in over twenty

years, removal of Broca's area produced fluent speech that
remained.
 Is Broca's area the speech center? Pribram went on to
argue that since its removal does not impair speech and may
even improve it, it is not necessary for speech production.
Nonetheless, it is possible that an intact and malfunctioning
Broca's area may interfere with the motor production of speech.
This, however, is a bit different from the conventional definition of
"speech center."

Penfield's "memories" suffered a related fate.

Penfield's Memories

Wilder Penfield was a Canadian neurosurgeon who operated on a

variety of patients, including epileptics. In many cases of epilepsy, anti-
epileptic drugs are ineffective and the sufferer is left undergoing
embarrassing and life-threatening seizures, with no recourse other than
a dangerous operation as a remedy.

The operation includes the removal of portions of cerebral cortex

that EEG analysis has suggested are responsible for the seizures. In
the course of the operation the patient is awake, while the
neurosurgeon probes for the offending tissue. If signs of a seizure can
be evoked, the cortical area is destroyed (e.g., by burning it with DC
through the electrode or by aspiration).
 In the course of these procedures in the late 1950s, Penfield found
that his stimulations, especially in the parietal and temporal cortex, often
aroused vivid memories in his patients, complete with color and sound.

Often an incident from many years ago seemed to be relived,

described by the patient as though it were being played on a
videocassette.

Penfield's discovery was described in countless introductory

psychology books and it seemed to add to the mounting evidence that the
phrenologists, though wrong in detail, were correct in general. The brain
seemed to be composed of many parts and each part has an obvious and
unique function. Penfield found stored memories, or so it seemed. We will
return to this issue.

What do you think was going on?

Lashley and the Engram
In a powerful display of faith in the precise localization of
function in the brain, a young Karl Lashley offered to trace the neural
connections in the frog brain and thus determine how the frog brain
worked. He had found some discarded slides of sections of the frog
brain and the neural connections seemed to him to be traceable. He
was shocked to learn that the stain used was extremely selective
and that the tissue he saw was therefore a very small fraction of the
total.
Despite his discouragement, he spent a substantial part of the
rest of his life trying to accomplish that mission. His findings had a
profound and lasting influence on the search for brain-behavior
relations Thompson & Robinson, 1978). In 1929 he published a
monograph, Brain Mechanisms and Intelligence, detailing his
findings concerning the effects of brain lesions on the ability of rats to
learn mazes.

What did he find?

 What he essentially found was that quite a bit of the cortex
had to be destroyed before any deficit was found and, surprisingly, that
it did not matter from what part of the cortex tissue was destroyed.
Thus, such and such a deficit would be found on the most
difficult maze with destruction of 50% of the cortex. But it did not seem
to matter what 50% was destroyed, as long as the primary projection
areas were spared. A rat that had one large lesion comprising half its
cortex would perform similarly to a rat that had twenty small lesions
scattered over the cortex. What was important was only the
percentage destroyed, not its location.
This means that the cortex is equipotential; any part can
carry out the function of any other part, within limits. The degree of the
deficit did increase with the size of the lesion, however, which led
Lashley to propose the principle of mass action. This means that
cortical tissue may be equipotential, but that its efficiency depends on
the amount which is remaining - the mass of cortex available is
important.
 The 1929 report was very discouraging to researchers aiming to
show localization of function. in the brain. If the particular locus of brain
damage is not crucial, then how can specific memories be stored in
specific places? Whether the memory trace (or engram) consists of
neural circuits, concentrations of proteins such as RNA, or presence of
neurotransmitters, the location of the lesions should be very important.

Lashley continued his search for a great many years, a search that
he described in 1950 ("In Search of the Engram"). After his 1929
monograph he tried slicing the cortices of his subjects, so that their
brains resembled sliced hams, only to find no deficits in learning tasks.
He destroyed the linkage between the sensory and the motor areas and
even lesioned the cerebellum. The cerebellum influences motor
behavior and, since other lesions had little effect, maybe the engrams
required to learn mazes were stored their. But even those subjects,
whose movements were hampered and who crawled, rolled, and
squirmed along the alley, came to the choice points and rolled down the
correct alley.

How can that be?

 Lashley's Legacy
Other data corroborate Lashley's basic findings and two
authoritative reviewers (Pribram, 1971; Thompson & Robinson,
1977) agree that his findings were legitimate. Instances of serious
brain damage producing little deterioration in performance abound.
For example, Chow (1970) destroyed three-quarters of the visual
cortex of cats and at the same time cut more than three-quarters
through their optic nerves. Such an operation would reduce the
animal's vision to near nil, one would think, and it does cause
disruption of a previously-learned visual discrimination.
But the cats relearned the discrimination as quickly as they had
originally learned! Other data show the discrimination performance
of cats and other animals to be passable after even greater
destruction of the visual pathway. It seems that a few hundred visual
cortical cells is sufficient for the learning of fairly difficult visual
discriminations (Lashley, 1970).
Is Your Brain Really Necessary?
That was the title of a piece appearing in Science (Lewin, 1980).
It describes the findings of a British neurologist, John Lorber, at the
University of Sheffield in the United Kingdom. Lorber's research
involves hydrocephalics, whose brain ventricles accumulate an excess
of cerebrospinal fluid.

When this occurs in an infant or young child, the skull expands to

make room for the excess fluid. But in older children and adults, the
skull is not malleable and the fluid crushes the forebrain against the
inside of the skull. In many instances great brain damage occurs,
accompanied by grave disturbances in function. However, in a great
many cases, there is no obvious deficit, even though the brain damage
is extreme. As Lorber put it:
“There's a young student at this university, who has an IQ of 126, has
gained a first-class honors degree in mathematics, and is socially
completely normal. And yet the boy has virtually no brain...When we did a
brain scan on him, we saw that instead of the normal 4.5-centimeter
thickness of brain tissue between the ventricles and the cortical surface,
there was just a thin layer of mantle measuring a millimeter or so. His
cranium is filled mainly with cerebrospinal fluid.” (Lewin, 1980, p. 1232)

The eminent British neuroanatomist , Patrick Wall, at University

College, London, commented that "Scores of similar accounts litter the
medical literature, and they go back a long way." He praised Lorber for
compiling a remarkable set of data, rather than relying on mere anecdotal
accounts. Wall wondered how we may account for such findings.
How indeed may we account for them? If Lorber is right and if
Lashley's search means anything, then we must at least question the old
supposition that the cerebral cortex is the seat of all intelligent behavior
and particularly that it is the repository for precisely localized memories.

But what of Penfield's famous findings?

Penfield Reconsidered
Penfield was no doubt sincere in his belief that he had found the
anatomical substrate for memory, but Valenstein (1973) showed that more recent
evidence paints a somewhat different picture. Fedio and Van Buren (1971), at
what was then the National Institute for the Study of Neurological Disease and
Stroke, point out that many surgeons use precisely the procedure used by
Penfield, yet no one seems to have found revived memories as he reported. That
fact, along with the report by Mahl (1970) that such patients often do report seeing
flashes of light or hearing brief sounds, suggests another interpretation for
Penfield's finding.

Imagine yourself as a patient under the conditions experienced by his patients.

You are undergoing a brain operation and, while you were under anesthesia,
the surgeon has cut through the scalp, the skull, and the dura under the skull.
You are now sitting there awake and the surgeon is touching your brain with a
stimulating electrode! It is surely fair to say that you might be a little "on edge"
or "reactive" under such circumstances.
 Suppose now that during the stimulation you see a flash of light or
hear a sound, just as you might see flashes and hear "bells ring" when
something strikes you in the head. You say, "I heard something," and the
surgeon asks what it was. Was it like a train whistle? Yes, it was and you
add, "I can see the station and there is my mother..." and so on. With a little
prodding, completely inadvertently done by the surgeon, a patient may well tell
many stories under such circumstances. Does this amount to the revival of
memories? In a sense it does, but the stimulation of the brain surface appears
to arouse only light flashes and brief auditory sensations. The elaborations of
these are aroused by the questions of the surgeon in the context of a highly
reactive subject.
Penfield was entirely well meaning and no reader of his biography,
No Man Alone (1970) could believe that he was intentionally perpetuating a
fraud. He found what his education had led him to expect and one cannot
blame him for believing that he found it. But others did not find it and the
reason for their failure is clear.
 Motivation and Emotion

“Thou shalt not sit

With statisticians nor commit
A social science”
(Auden, 1946, st. 27)

Probably the best known discoveries in physiological psychology occurred

in the area of motivation and emotion; this includes the discovery of what some
called "reward" and "punishment" centers, feeding centers, aggression centers,
and the like. The story involves the limbic system, particularly the
hypothalamus.

******[DRAW DIAGRAM]******
If the brain were an apple, its core would be the limbic system, old cortex
arranged essentially the same in us as it is in dogs, rabbits, and rats. Limbic
means "border" and refers to the brain tissue bordering the midline of the brain.
The hypothalamus, a cluster of cell bodies about as large as the tip of your
thumb, is a crucial part of this system and controls the autonomic nervous
system.
 The autonomic nervous system is subdivided into the sympathetic
and the parasympathetic branches. The division was suggested by two
Viennese neurologists, Karplus and Kreidl, in 1909 and subsequent
research has supported their view. They suggested that the anterior
(forward) portion of the hypothalamus controls parasympathetic activity;
this includes conservative functions such as sleep, sexual activity,
feeding, and other "vegetative" functions.

On the other hand, the posterior (rearward) hypothalamus controls

aggression, flight, and other activities that use up energy in the
interests of survival. The posterior hypothalamus increases
sympathetic activity, which means that heart rate increases, blood is
shunted from the viscera to the muscles, the lungs exchange gases
more rapidly, and the affected organism is more able to fight for its life
or to flee. And Karplus and Kreidl were correct, as many studies since
have shown. The hypothalamus seems to be a center for motivation
and emotion and in a sense it is.
Papez's Circuit
In 1936 a neuroanatomist, James Papez, made a bizarre proposal, based
wholly on anatomical evidence and reasonable assump-tions about the nervous
system and emotion.

[EXPLAIN]
We now know that stimulation of the brainstem reticular formation (Moruzzi &
Magoun, 1949; Hebb, 1954) produces nonspecific arousal of the entire forebrain.

[EXPLAIN]

In 1942 Hetherington and Ranson found what came to be called the "satiety
center.“

[EXPLAIN]
 In 1951 Anand and Brobeck at Yale University found the "feeding
center." While attempting to insert stimulating electrodes into the amygdala
of rats (the amygdala is covered below), they inadvertently destroyed the
lateral nuclei of the hypothalamus and found that their subjects died after the
operation. The cause of death was aphagia; the rats refused to eat and spat
out food that was forced in their mouths.

Thus, the hypothalamus seemed to control both eating and not

eating. If the VM "satiety" center were damaged, hyperphagia, or gross
overeating, occurred. If the lateral nuclei (LH) were destroyed, eating
ceased. Reasonably enough, when the VM was stimulated, eating ceased,
and stimulation of the LH produced eating. Here were the feeding and
satiety centers and everyone accepted them. The only question was what
new centers would be discovered to account for new behaviors? No wonder
Hess won the Nobel Prize.
Re-evaluation of LH and VMH Function

A great deal of evidence indicates that the effects of VM and LH lesions

are more general than had been believed and that the ordinary functions of
the two structures are inhibitory and excitatory, respectively. Destruction of
the VM removes an inhibitory influence and leads to increased
responsiveness to strong external cues.
The LH appears excitatory, so that stimulation of it appears in some
ways to act as does destruction of the VM. Such stimulation can maintain
lever pressing, produce feeding, cause stalking of prey, and other activities
that seem describable as "outer directed." Destruction of the LH causes
decreased responsiveness to external stimuli and even to one's own body.
Operated cats will remain immobile, even when placed in uncomfortable
positions, and will ignore stimuli that would ordinarily elicit strong reactions.

This sensory neglect is a powerful general effect, yet, as Carlson (1991)

noted, the lateral hypothalamus was called the "feeding center" for
approximately two decades! Therein lies a lesson for us.
 Add to This List (see MIT Press History Chapter 10 online at the Nuoro
page of www.geocities.com/malonejc2007. There is far more detail than
will appear in the actual book.)

The Amygdala as “aggression center”

Kluver/Bucy Effect and psychosurgery
Effects of bilateral amygdalectomy
Freeman & Watts
Downer’s finding
James Olds and “pleasure centers”
Hebb
Delgado
Valenstein
• Broca’s “Language Center” (motor control)
• Penfield’s Memories (surgeon’s prompting)
• Morruzi & Magoun’s “Arousal Center” (EEG only)
• “Satiety and Feeding Centers” (E/I control)
• “Aggression Centers” (psychic blindness)
• “Pleasure/Reward Centers” (also E/I control)
In all of these cases, misinterpretations of effects were
made and, much later, a better-informed interpretation
arose.
So…..beware of claims of “breakthroughs!”